CN103710357B - Alfalfa stress response gene M sNAC2 and application thereof - Google Patents
Alfalfa stress response gene M sNAC2 and application thereof Download PDFInfo
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- CN103710357B CN103710357B CN201310717761.4A CN201310717761A CN103710357B CN 103710357 B CN103710357 B CN 103710357B CN 201310717761 A CN201310717761 A CN 201310717761A CN 103710357 B CN103710357 B CN 103710357B
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Abstract
The present invention relates to a kind of alfalfa stress response gene M sNAC2 and application thereof.The present invention's application RT-PCR and RACE technology clone NAC family genes involved from alfalfa genome; called after MsNAC2; its nucleotide sequence as shown in SEQ ID No.11, and constructs the fusion expression vector containing reinforced green fluorescent protein (EGFP) gene.The present invention's application fluorescent quantitative PCR technique analyzes the expression pattern of this gene in alfalfa, and this gene and environment stress (damage to plants caused by sudden drop in temperature, salt damage, arid) between relation, find that this gene process LAN in tobacco can improve cold-resistant, drought resisting and the salt tolerance of transgenic tobacco plant, this gene can be used for the genetic transformation of other single dicotyledonss, improves its resistance.
Description
Technical field
The invention belongs to genetically engineered field, specifically, relate to alfalfa stress response gene M sNAC2 and application thereof.
Background technology
NAC transcription factor (comprising NAM, ATAF1/2 and CUC2) is the distinctive a kind of transcription factor with multiple regulating and controlling effect of plant, in extensive existence and terrestrial plant, the main structure of this kind of transcription factor is the NAC structural domain that N holds as conservative about 150 amino-acid residues, can, in conjunction with DNA and other albumen, can be divided into from A ~ E5 conserved regions; C end is then the transcriptional activation control region of great diversity, is rich in Serine, Threonine, proline(Pro), L-glutamic acid etc., and research finds that the C-end of some NAC albumen has protein binding activity.There are some researches show that some members of NAC family are worked in Stress response network at present.In Arabidopis thaliana, ANAC019, ANAC055 and ANAC072 are subject to the abduction delivering of arid, high salt and dormin (ABA), and they can significantly improve the drought-resistance ability (Tran et al., 2004) of plant.Arabidopis thaliana LOV1 gene can strengthen the freezing tolerance (Yoo et al., 2007) of plant.SNAC1/2 (stress-responsive NAC2) gene from paddy rice is subject to the abduction delivering of arid, high salt, low temperature, wound and ABA, and cold-resistant, the anti-salt of its process LAN plant also can resist arid (Hu et al., 2008).Although the paddy rice of overexpression OsNAC6 gene grows slowly, yields poorly, demonstrate the stronger resistance (Nakashimaet al., 2007) to arid, high salt and blight.Although the function of NAC gene need further investigation, they have shown huge application potential in genetically engineered plant.
Summary of the invention
The object of this invention is to provide alfalfa stress response gene M sNAC2 and application thereof.
The invention provides alfalfa stress response gene M sNAC2, it has:
1) nucleotide sequence shown in SEQ ID No.11; Or
2) shown in SEQ ID No.11, nucleotide sequence is substituted, lacks and/or increases one or several Nucleotide; Or
3) under strict conditions with 1) DNA sequence dna that the limits nucleotide sequence of hybridizing.
The invention provides the expression vector containing said gene MsNAC2.
In one embodiment of the invention, the expression vector containing gene M sNAC2 obtained is pBIGFP-MsNAC2.
The construction process of this expression vector is: the cDNA reversed with the total serum IgE of alfalfa-leaf is template, carries out PCR reaction, and primer sequence is:
Upstream primer: 5 '-TCTAGAATGATCAGTCGATGGAAACACTTC-' 3(SEQ ID NO.13)
Downstream primer: 5 '-GGTACCGACTTGGGCAGATACATAAAGAC-' 3(SEQ ID NO.14);
Be connected on pMD18-T carrier after RT-PCR amplified production is reclaimed, connect product conversion e.colistraindh5α, choose positive colony to check order, with XbaI and KpnI, MsNAC2 gene is got off from unloading pMD18-T, be connected in XbaI and the KpnI site containing eGFP plant expression vector pBIGFP after recovery, build and obtain pBIGFP-MsNAC2.
Host cell containing above-mentioned expression vector also belongs to protection scope of the present invention.
Present invention also offers the transformed plant cells containing gene M sNAC2.
The invention provides the application of MsNAC2 gene in preparation transgenic plant.
Preferably, described plant is Arabidopis thaliana, tobacco, onion, paddy rice or cotton.
The invention provides MsNAC2 gene and improve the application in stress resistance of plant.
Described resistance refers to cold-resistant, drought resisting or salt tolerance.
Present invention also offers the method for clone's alfalfa stress response gene M sNAC2, comprise the following steps:
(1) utilize RT-PCR method to increase from alfalfa genome and obtain the conserved regions fragment sequence of NAC;
(2) according to the conserved regions sequences Design 3 ' RACE primer of step (1), then with alfalfa total serum IgE for template, carry out 3 ' RACE fragment amplification;
(3) 5 ' RACE primer amplification 5 ' RACE fragment is designed;
(4) use DNAMAN5.2 software by conserved regions sequence, 3 ' RACE fragment becomes complete complete genome sequence with 5 ' RACE fragment assembly.
Wherein, in the RT-PCR method of step (1), the primer sequence is:
Upstream primer sequence is 5 '-ACCAAACGGTTCAAGGCCGAACC-' 3(SEQID NO:1), downstream sequence is 5 '-CGATACTCGTGCATGATCCAATTG-' 3(SEQ ID NO:2).
The total serum IgE of step (1) alfalfa obtains cDNA after reverse transcription, and its PCR response procedures is: 94 DEG C of denaturation 5min, (94 DEG C of 10s, 55 DEG C of 20s, 72 DEG C of 1min) 30 circulation, 72 DEG C of 10min.
Wherein, 3 ' RACE primer sequence of step (2) is 5 '-GTTCAAGGCCGAACCGGGCTG-3 ' (SEQ ID NO:3).This step uses 3 '-Full RACE Set Agarose Regular test kit amplification 3 ' RACE fragment.
Wherein, the 5 ' RACE primer 5 '-AGCCCGGTTCGGCCTTGAACC-3 ' (SEQ ID NO:4) of step (3).This step uses 5 '-Full RACE Set Agarose Regular test kit amplification 5 ' RACE fragment.
The present invention has following advantages and beneficial effect:
The invention provides alfalfa stress response gene M sNAC2(nucleotide sequence as shown in SEQ ID No.11) and application in crop breeding.This gene not only by low temperature and high Salt treatment high expression level, but also is expressed by arid and ABA stress-inducing.The research of onion epidermis Subcellular Localization shows that MsNAC2 is nuclear location gene.The adversity gene of previous report functionally majority has unicity, and research finds that MsNAC2 gene of the present invention is improving in cold-resistant, the drought resisting of transgenic tobacco plant and salt tolerance etc. all according to there being obvious effect.This gene can be used for the genetic transformation of other single dicotyledonss, significantly improves its resistance.
Accompanying drawing explanation
Fig. 1 is fusion expression vector pBIGFP-MsNAC2 structural representation.
Fig. 2 is expression vector pBIGFP structural representation.
Fig. 3 is the relative expression of MsNAC2 after stress-inducing.Wherein, A figure is the relative expression of MsNAC2 in blade and root under 250mMNaCl Stress treatment; B figure is the relative expression of MsNAC2 in blade and root under 10%PEG6000 Stress treatment; C figure is 100 μMs of lower MsNAC2 relative expressions in blade and root of ABA induction; D figure is the lower relative expressions of MsNAC2 in blade and root of 4 DEG C of process.
Fig. 4 is MsNAC2 gene Subcellular Localization photo.Wherein, A figure is the picture of transgene tobacco under white light containing MsNAC2-GFP fusion gene; B figure is the transgene tobacco picture under ultraviolet light containing MsNAC2-GFP fusion gene; C figure is that control group is only containing the picture of transgene tobacco under white light of GFP green fluorescence protein gene; D figure is that control group is only containing the transgene tobacco picture under ultraviolet light of GFP green fluorescence protein gene.
Fig. 5 turns MsNAC2 genetic tobacco growing state under environment stress, and after 250mM NaCl process 20d, transgene tobacco grows fine, and wild-type tobacco blade almost whole chlorosis, become faint yellow.
Fig. 6 turns MsNAC2 genetic tobacco growing state under environment stress, and after 10%PEG process 20d, transgene tobacco grows fine, and wild-type tobacco blade almost all becomes water stain shape.
Fig. 7 turns MsNAC2 genetic tobacco growing state under environment stress, and after carrying out renewal cultivation 10d again after 4 DEG C of process 6d, transgenic tobacco plant blade obviously will be greater than wild-type, and root system is also flourishing than wild-type.
Embodiment
Following examples for illustration of the present invention, but are not used for limiting the scope of the invention.Without departing from the spirit and substance of the case in the present invention, the amendment do the inventive method, step or condition or replacement, all belong to scope of the present invention.
If do not specialize, the conventional means that technique means used in embodiment is well known to those skilled in the art; If do not specialize, in embodiment, agents useful for same is commercially available.
The acquisition of embodiment 1 alfalfa stress response gene M sNAC2
First according to leguminous plants peanut NAC transcription factor Gene A hNAC2 and AhNAC3(EU755023 and EU755022) sequence, design clover NAC DNA homolog primer, upstream primer sequence is 5 '-ACCAAACGGTTCAAG GCCGAACC-' 3(SEQ ID NO.1), downstream sequence is 5 '-CGATACTCGTGCATGATCCAATTG-' 3(SEQ ID NO.2).Then, utilize RT-PCR technology first to increase from alfalfa genome and obtain the conserved regions fragment sequence of NAC.
1, the extraction of alfalfa total serum IgE: adopt TaKaRa RNAiso Plus to extract total serum IgE.
2, the synthesis of cDNA first chain:
Reaction system is as follows:
Reverse transcription program is: 42 DEG C of 60min, 70 DEG C of 15min.
3, PCR reaction:
PCR response procedures is: 94 DEG C of denaturation 5min, (94 DEG C of 10s, 55 DEG C of 20s, 72 DEG C of 1min) 30 circulation, 72 DEG C of 10min.
4, conserved regions clone: above-mentioned RT-PCR amplified production Takara DNA gel is reclaimed test kit and reclaims, reclaim product and be connected on pMD18-T carrier, operation steps is undertaken by Takara pMD18-T specification sheets.Connect product conversion e.colistraindh5α, LB substratum containing 100mg/L penbritin is cultivated, picking mono-clonal bacterium colony, plasmid is extracted with Takara plasmid extraction kit, after the qualification of EcoRI and Hind III double digestion, choose positive colony to check order, checking order is completed by the handsome biotech firm in Beijing.
5、RACE
According to conserved regions primers:
3 ' RACE upstream primer: 5 '-GTTCAAGGCCGAACCGGGCTG-3 ' (SEQ ID NO.3)
5 ' RACE downstream primer: 5 '-AGCCCGGTTCGGCCTTGAACC-3 ' (SEQ ID NO.4)
3 '-Full RACE Set Agarose Regular test kit is used to carry out 3 ' RACE fragment amplification.5 '-Full RACE Set Agarose Regular test kit is used to carry out 5 ' RACE fragment amplification.
6, finally use DNAMAN5.2 software by conserved regions sequence, 3 ' RACE fragment becomes complete complete genome sequence with 5 ' RACE fragment assembly.Splicing length is 1358bp, and nucleotide sequence is as shown in SEQ ID NO.11, and the aminoacid sequence of coding is as shown in SEQ ID NO.12.Find in this sequence inputting NCBI blastp, aminoacid sequence and the leguminous plants NAC family gene sibship of this genes encoding are very near.Be such as 89% with M. truncatula NAC genetic homology, being 87% with garbanzo NAC genetic homology, is structural homology > 77% with soybean NAC genetic homology.Therefore this gene is the gene of the NAC family that is newly identified, and by its called after MsNAC2.
Embodiment 2 real-time fluorescence quantitative PCR detects the expression characterization of MsNAC2 gene under environment stress condition in alfalfa
The growth seedling of alfalfa of two weeks is proceeded to respectively containing 250mmolL
-1naCl, 10%PEG6000,100 μm of olL
-1carry out Stress treatment in dormin (ABA) MS liquid nutrient medium, damaging to plants caused by sudden drop in temperature process is then carry out in 4 DEG C of refrigerators.Treatment time of PEG6000, ABA and 4 DEG C is 1,2,4,8,12 and 24h, treatment time of NaCl is 1,2,4,8,12,24 and 48h, with contrasting of not dealing with, process the rear total serum IgE extracting blade and root respectively, full length cDNA sequence design fluorescent quantitation according to MsNAC2 expresses primer, upstream primer sequence is 5-TGGTGAAGATGACCCTTTTGC-3 ' (SEQ IDNO:5), and downstream primer sequence is 5 '-AAGCTCCACTTGCAGTTGCAG-3 ' (SEQID NO:6).With alfalfa Actin gene (EU664318) for reference gene, reference gene upstream primer sequence is 5 '-CAGGTCGTGATCTCACAGACG-3 ' (SEQ IDNO:7), and downstream primer sequence is 5 '-TCTTCTCAACAGCTGAGCTCG-3 ' (SEQID NO:8).Real-time Real-Time PCR quantitative analysis is carried out, see Fig. 3 with the sample of SYBR Green I method to Different stress treatment time point.Within the treatment time of this experiment setting, MsNAC2 coerces lower inducing effect the most obviously (the D figure in Fig. 3) cold, and expression amount during 2h in root reaches climax, for 4.408 times of contrast, expression amount reduces subsequently, reaches second peak value when 24h, is 3.387 times of contrast.Expression in blade first reduces then to raise gradually, is 1.029 times of contrast when 12h, is 2.412 times of contrast during 24h; PEG coerces down (the B figure in Fig. 3), and during 1h, MsNAC2 expression amount reaches first peak value in root, and be 1.496 times of contrast, expression amount reduces subsequently, reaches second peak value when 12h, is 1.347 times of contrast.And the expression amount in blade is all lower than control group; Under ABA induction (the C figure in Fig. 3), during 1h, MsNAC2 expression amount reaches climax in root, is 1.424 times of contrast, reduces gradually subsequently.Expression amount in blade is equally all lower than control group; The expression trend that MsNAC2 presents under 250mM NaCl coerces first declines to then raising, and when 48h, MsNAC2 expression amount is 1.469 times of contrast in root.The expression amount of same MsNAC2 gene in root will apparently higher than the expression amount (the A figure see in Fig. 3) in blade.Quantitative analysis results shows, this gene damage to plants caused by sudden drop in temperature coerce under in root and blade equal up-regulated expression, salt stress, drought stress and ABA induction under, at root up-regulated expression, in blade expression amount lower.
The Subcellular Localization of embodiment 3MsNAC2 gene
By gene constructed for MsNAC2 in the fusion expression vector pBIGFP containing eGFP gene, be built into fusion expression vector pBIGFP-MsNAC2(detailed in Example 4).Then in conjunction with agriculture bacillus mediated transient expression technology and laser co-focusing detection technique, successfully expression vector pBIGFP-MsNAC2 is transformed onion epidermis.Then at the expressive site of this gene of fluorescence microscopy Microscopic observation, not contain the carrier pBIGFP of MsNAC2 for contrast, the results are shown in Figure 4, wherein, turn the onion epidermis cell of pBIGFP-MsNAC2, only can observe GFP green florescent signal (the B figure in Fig. 4) under ultraviolet light in nucleus, the A figure in Fig. 4 is the photo of onion epidermis cell under white light turning pBIGFP-MsNAC2; And control group, can observe the green florescent signal (the D figure in Fig. 4) of GFP clearly in the nucleus turning the onion epidermis cell of pBIGFP and in tenuigenin, the C figure in Fig. 4 is the photo of onion epidermis cell under white light turning pBIGFP.Result shows that MsNAC2 gene is nuclear location gene.
The structure of embodiment 4 plant expression vector pBIGFP-MsNAC2
1, primer is designed according to isolated alfalfa stress response gene M sNAC2:
Upstream primer: 5 '-TCTAGAATGATCAGTCGATGGAAACACTTC-' 3(SEQ ID NO.13)
Downstream primer: 5 '-GGTACCGACTTGGGCAGATACATAAAGAC-' 3(SEQ ID NO.14)
The cDNA reversed with the total serum IgE of alfalfa-leaf is template, carries out PCR reaction.
2, above-mentioned RT-PCR amplified production is reclaimed, reclaim product and be connected on pMD18-T carrier, connect product conversion e.colistraindh5α, choose positive colony and check order.
3, got off from unloading pMD18-T by MsNAC2 gene with XbaI and KpnI, be connected to after recovery in XbaI and the KpnI site containing eGFP plant expression vector pBIGFP, plant expression vector construction completes, called after pBIGFP-MsNAC2.This carrier structure figure is shown in Fig. 1.
Embodiment 5 alfalfa stress response gene M sNAC2 transformation of tobacco
By the plant expression vector transformation Agrobacterium LBA4404 built.With electric shocking method, plasmid pBIGFP-MsNAC2 is forwarded in Agrobacterium LBA4404, then tobacco leaf disc is infected with Agrobacterium, leaf dish is under the selection containing kantlex screening culture medium (MS+0.5mg/L BA+0.01mg/LIAA+Kana50mg/L), successful induction and filter out resistance budlet, resistance budlet is proceeded in root media (1/2MS+0.5mg/L IBA+Kana50mg/L) and induce healthy and strong root, then move into hardening in flowerpot, obtain transgene tobacco.
Random selecting 10 strain transgene tobacco, extracts DNA and carries out PCR detection, take wild-type tobacco as contrast.PCR detects, primer is designed according to the resistance screening gene NPTII in parallel with goal gene MsNAC2, upstream primer is 5'-AACAGACAATCGGCTGCTCT-3'(SEQ ID NO.9), downstream primer is 5'-CCACCATGATATTCGGCA AGCA-3'(SEQ ID NO.10), PCR can amplify the fragment of 550bp.PCR result shows, this 10 strain transformed plant is all PCR positive material, and this preliminary proof foreign gene has been incorporated in tobacco gene group.(NPTII gene and MsNAC2 gene are connected in parallel on same expression vector, and NPTII is a kind of marker gene on pBIGFP-MsNAC2 carrier)
The functional analysis of embodiment 6 alfalfa stress response gene M sNAC2 transformation of tobacco positive plant
After the positive transgenic Tobacco Flowering that example 5 to be performed obtains is solid, collects T1 and carry out stress experiment for transgenic seed.
Transgenic Tobacco Seeds and wild-type tobacco seed are sowed at MS simultaneously
0substratum germinates, sprouts after 20d until young shoot, young shoot is transplanted respectively to respectively containing in the MS substratum of 250mMNaCl and 10%PEG, after 20d, record result.When damaging to plants caused by sudden drop in temperature process, the tobacco seedling of the rear 30d that germinates is placed in 4 DEG C of refrigerators, carries out Recovery processing 10d again after deepfreeze 6d, then record result.
(1), after 250mM NaCl process 20d, transgene tobacco grows fine, and wild-type tobacco blade almost whole chlorosis, become faint yellow, the results are shown in Figure 5.(salt tolerance)
(2), after 10%PEG process 20d, transgene tobacco grows fine, and wild-type tobacco blade almost all becomes water stain shape, the results are shown in Figure 6.(drought resistance)
After carrying out renewal cultivation 10d again after (3) 4 DEG C of process 6d, transgenic tobacco plant blade obviously will be greater than wild-type, and root system is also flourishing than wild-type, the results are shown in Figure 7.(cold resistance)
The above results proves, deriving from alfalfa NAC transcription factor gene (MsNAC2) is an effective adverse circumstance genes involved, can be applied in the operation of cultivating transgenosis adversity resistant plant, can significantly improve the resistance of plant.
The above is only the preferred embodiment of the present invention; it should be pointed out that for those skilled in the art, under the prerequisite not departing from the technology of the present invention principle; can also make some improvements and modifications, these improvements and modifications also should be considered as protection scope of the present invention.
Claims (3)
1. the application in the transgene tobacco that improves in preparation resistance of alfalfa stress response gene, described alfalfa stress response gene is MsNAC2, and nucleotides sequence is classified as shown in SEQ ID No.11, and described resistance is cold-resistant, drought resisting or salt tolerance.
2. the application of the expression vector containing alfalfa stress response gene in the transgene tobacco of preparation resistance raising, it is characterized in that, described expression vector is pBIGFP-MsNAC2, described alfalfa stress response gene is MsNAC2, nucleotides sequence is classified as shown in SEQ ID No.11, and described resistance is cold-resistant, drought resisting or salt tolerance.
3. the application of host cell in the transgene tobacco of preparation resistance raising, it is characterized in that, described host cell contains the expression vector of the alfalfa stress response gene that can improve stress resistance of plant, described expression vector is pBIGFP-MsNAC2, described alfalfa stress response gene is MsNAC2, nucleotides sequence is classified as shown in SEQ ID No.11, and described resistance is cold-resistant, drought resisting or salt tolerance.
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| CN106967728B (en) * | 2017-04-13 | 2020-04-24 | 华中农业大学 | Pumpkin stress resistance gene CmNAC1 and application thereof |
| CN108486129A (en) * | 2018-02-27 | 2018-09-04 | 中国农业大学 | Plant frigostabile Gene A tBTF3L and its application |
| CN113308479B (en) * | 2021-07-15 | 2022-04-29 | 浙江大学 | Application of SlNAC100 gene in improvement of low-temperature resistance of tomato |
| CN116003546B (en) * | 2022-07-18 | 2024-03-08 | 上海交通大学 | Alfalfa NAC transcription factor and application thereof |
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